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Biomedical subjects

C G Farmer

Publications and source records attributed to C G Farmer.

At least 19 recordsLinked to original sources

On the origin of avian air sacs.

For many vertebrates the lung is the largest and lightest organ in the body cavity and for these reasons can greatly affect an organism's shape, density, and its distribution of mass; characters that are important to locomotion. In this paper non-respiratory functions of the lung are considered along with data on the respiratory capacities and gas exchange abilities of birds and crocodilians to infer the evolutionary history of the respiratory systems of dinosaurs, including birds. From a quadrupedal ancestry theropod dinosaurs evolved a bipedal posture. Bipedalism is an impressive balancing act, especially for tall animals with massive heads. During this transition selection for good balance and agility may have helped shape pulmonary morphology. Respiratory adaptations arising for bipedalism are suggested to include a reduction in costal ventilation and the use of cuirassal ventilation with a caudad expansion of the lung into the dorsal abdominal cavity. The evolution of volant animals from bipeds required yet again a major reorganization in body form. With this transition avian air sacs may have been favored because they enhanced balance and agility in flight. Finally, I propose that these hypotheses can be tested by examining the importance of the air sacs to balance and agility in extant animals and that these data will enhance our understanding of the evolution of the respiratory system in archosaurs.

Air Sacs↗

Contribution of the diaphragmaticus muscle to vital capacity in fasting and post-prandial American alligators (Alligator mississippiensis).

The importance of the diaphragmaticus muscle to vital capacity was investigated in juvenile American alligators by transection of this muscle. In both fasting and post-prandial animals a pneumotach was used to study vital capacity that was stimulated by either a hypercapnic-anoxic gas mixture or a hypercapnic-normoxic gas mixture in two types of control groups of animals (a shamoperated group and a group receiving no treatment) and in the experimental (transected) group. Transection did not significantly reduce vital capacity or affect time of inspiration or expiration in fasted animals. For both the experimental and control groups vital capacity was greatly reduced in post-prandial animals compared to the fasting state. Furthermore, alligators with a transected diaphragmaticus muscle showed a 16-18% greater drop in vital capacity in the post-prandial state than did alligators with an intact diaphragmaticus muscle. The post-prandial decrease in vital capacity for alligators with a transected diaphragmaticus occurred concomitantly with a significant increase in time to inspire and a decrease in maximum rate of inspiration when compared to control animals. The results from this study suggest that the diaphragmaticus muscle plays an important role in enabling large volumes of oxygen to be taken into the lungs in the post-prandial state.

Alligators and Crocodiles↗

Structure and function of the esophagus of the American alligator (Alligator mississippiensis).

Esophageal structure and function were studied in juvenile American alligators (Alligator mississippiensis). The anatomy of alligators differs from humans in several important aspects: the crocodilian esophagus is more muscular and is composed entirely of smooth muscle. Functionally, the crocodilian esophagus is similar to that of mammals, but alligators have peak esophageal peristaltic pressures that are 2-3-fold greater than pressures in the human esophagus. As is found in humans, the incidence of esophageal reflux increased in postprandial animals compared with the fasting state. We observed a large increase in pressure in the lower esophageal sphincter (LES) during ventilation that ranged from 200% to 3000% of the pressures measured during apnea. These pressure changes appear to be intrinsic to the LES. Alligators lack a mammalian-type diaphragm; thus, there is no crural diaphragmatic contribution to LES pressure. These features recommend the alligator as a useful model for the study of regulation of the LES.

Alligators and Crocodiles↗

Effect of supplementation frequency and supplemental urea level on dormant tallgrass-prairie hay intake and digestion by beef steers and prepartum performance of beef cows grazing dormant tallgrass-prairie.

Effect of supplementation frequency and supplemental urea level on forage use (Exp. 1) and performance (Exp. 2 and 3) of beef cattle consuming low-quality tallgrass-prairie were evaluated. For Exp. 1 and 2, a 2 x 2 factorial treatment structure was used, such that two supplements (30% CP) containing 0 or 30% of supplemental degradable intake protein (DIP) from urea were fed daily or on alternate days. In Exp. 1 and 2, supplement was fed at 0.41% BW daily or at 0.83% BW (DM basis) on alternate days. For Exp. 3, a 2 x 4 factorial treatment structure was used, such that four supplements (40% CP) containing 0, 15, 30, or 45% of supplemental DIP from urea were fed daily or 3 d/wk. Supplements were group-fed at 0.32% BW daily or at 0.73% BW (DM basis) 3 d/wk. In Exp. 1, 16 Angus x Hereford steers (initial BW = 252 kg) were blocked by BW and assigned to treatment. Urea level x supplementation frequency interactions were not evident for forage intake, digestion, or rate of passage. Forage OM intake (OMI) and total digestible OMI (TDOMI) were not significantly affected by treatment. Total-tract digestion of OM (P = 0.03) and NDF (P = 0.06) were greater for steers supplemented daily. In Exp. 2, 48 Angus x Hereford cows (initial BW = 490 kg) grazing winter tallgrass prairie were used. Significant frequency x urea interactions were not evident for BW and body condition (BC) change; similarly, the main effects were not substantive for these variables. In Exp. 3, 160 Angus x Hereford cows (initial BW = 525 kg) grazing dormant, tallgrass prairie were used. Supplement refusal occurred for cows fed the highest urea levels, particularly for cows fed the supplement with 45% of the DIP from urea 3 d/wk, and supplement refusal increased closer to calving. A frequency x urea interaction (P = 0.02) was observed for prepartum BW changes. As supplemental urea level increased, prepartum BW loss increased quadratically (P = 0.02); however, a greater magnitude of loss occurred when feeding supplements containing > or = 30% of DIP from urea 3 d/ wk. Cumulative BC change followed a similar trend. In conclusion, moderate protein (< or = 30% CP) supplements with < or = 30% of supplemental DIP from urea can be fed on alternate days without a substantive performance penalty. However, infrequent feeding of higher protein (> 30% CP) supplements with significant urea levels (> 15% of DIP from urea) may result in decreased performance compared with lower urea levels.

Animal Feed↗

Ruminal and host adaptations to changes in frequency of protein supplementation.

The effect of altering supplementation frequency on host N balance and key N transactions in the ruminal ecosystem were monitored. Four ruminally fistulated beef steers (BW = 513 kg; SEM = 6.5) were used in a 2 x 2 crossover design with two periods and two supplementation frequency treatments. Supplementation frequencies were 2 and 7 d/wk. Steers were fed tallgrass prairie hay (73.1% NDF, 5.3% CP) ad libitum. Supplement (42% CP; DM basis) was fed at 0.36% BW/d to steers supplemented 7 d/wk. Steers supplemented 2 d/wk received the same amount of supplement per week, but it was equally split among the two supplementation events. Steers supplemented 7 d/wk had higher forage (P < 0.02) and total digestible OM intake (P < 0.06), total N intake, fecal N excretion, and N retention. Although both supplementation frequencies were characterized by positive N balance, the decrease in N retention in the steers supplemented 2 d/wk was due to higher (P < 0.01) urinary N loss. Ruminal fluid was sampled at 0, 2, 4, 6, 12, 24, 48, and 72 h after supplementation beginning on a day when both treatments were supplemented. Frequency x hour interactions (P < 0.02) were observed for ruminal N metabolism criteria. Counts of peptide- and AA-fermenting bacteria peaked at 2 h and returned to nadir by 12 h for steers supplemented 7 d/wk. Steers supplemented 2 d/wk peaked at 6 h with a greater population and returned to nadir at 48 h. Ruminal ammonia concentrations followed a similar trend. Specific activity of ammonia production was lower (P < or = 0.05) immediately after supplementation for steers supplemented 2 d/wk, but by 12 h was the same as for 7 d/wk steers. Ruminal peptides and free AA peaked at 2 h for steers supplemented 2 d/wk and were generally higher (P < or = 0.05) during the first 6 h compared with steers supplemented 7 d/wk. Total VFA concentration was not different (P = 0.35) due to supplementation frequency. Frequency x hour interactions (P < 0.01) were observed for all molar proportions of VFA. The molar proportion of acetate and acetate:propionate ratio were lower (P < 0.01) and the molar proportions of propionate and butyrate were higher for steers supplemented 2 d/wk from 4 h to 24 h. In conclusion, forage use and N balance improved with supplementation 7 d/wk, but supplementation 2 d/wk was associated with some desirable shifts in select ruminal events that may contribute to moderating potential negative impacts of supplementing infrequently.

Adaptation, Physiological↗

Effect of ruminal vs postruminal administration of degradable protein on utilization of low-quality forage by beef steers.

An experiment was designed to determine the effects of ruminal and postruminal infusions of ruminally degradable protein (casein) on intake and digestion of low-quality hay by beef steers. Twelve ruminally fistulated Angus x Hereford steers (initial BW = 563 kg) were blocked by weight and assigned to one of three treatments: control (C; hay only) or hay plus ruminal (R) or postruminal (P) infusion of 400 g/d of sodium caseinate. The trial consisted of five periods: 1) 10-d adaptation to the hay diet; 2) 7-d measurement of hay intake (without infusions); 3) 10-d adaptation to protein infusion treatments (intake measurements continued); 4) 7-d measurement of hay intake and digestibility (infusions continued); and 5) 3-d ruminal sampling period (infusions continued). Steers were given ad libitum access to tallgrass-prairie hay (3.4% CP, 76.6% NDF) throughout the study. Casein was administered once daily before feeding, either directly into the rumen or via anchored infusion lines into the abomasum. Hay intake was increased by supplementation (P < 0.01). Ruminal infusion elicited a greater (P = 0.04) increase in hay intake than postruminal infusion. Intake tended (P = 0.11) to be lower in period 4 than in period 2 for control steers but was greater in period 4 than in period 2 (P < or = 0.03) for both R and P steers. The increase in intake between periods 2 and 4 was greater for R than for P steers (P = 0.03). Supplementation improved diet OM digestion (P = 0.04) but not NDF digestion (P = 0.18); however, greater relative error for NDF digestion may have limited the ability to elucidate significant treatment effects. There were no differences in either OM digestion (P = 0.42) or NDF digestion (P = 0.35) between R and P steers. Plasma urea N at 0 and 3 h after feeding on the last day of the experiment was lower (P = 0.05) for C than for R and P steers, but no difference (P = 0.48) was evident between R and P steers. Ruminal ammonia N levels also were increased by supplementation (P < 0.01), with a much larger increase for R than for P steers (P < 0.01). Total VFA concentrations were not affected (P = 0.21) by treatment, but R steers exhibited lower proportions of acetate and higher proportions of isobutyrate, valerate, and isovalerate than P steers (P < 0.01). In conclusion, ruminal and postruminal infusion of a degradable protein source improved forage utilization, although the response in forage OM intake and total digestible OM intake was greater for ruminal infusion than for postruminal infusion.

Animal Feed↗

The effects of several supplementation frequencies on forage use and the performance of beef cattle consuming dormant tallgrass prairie forage.

Two experiments were conducted to quantify the impact on forage use and performance of varying supplementation frequency of cattle consuming forage diets across a range of frequencies. In both experiments, a common supplement was used that contained a relatively high concentration of CP (43%) and was fed at the following frequencies: 1) 2 d/wk; 2) 3 d/wk; 3) 5 d/wk; and 4) 7 d/wk. In Exp. 1, 120 Hereford x Angus cows (BW = 537 kg) grazing winter tallgrass-prairie range were supplemented at the various frequencies from December 7 until calving (average calving date = 3/7/99). All treatments provided the same quantity of supplement on a weekly basis (12.74 kg, as-fed) but divided the amount delivered on a given day equally among the number of supplementation events for that treatment. Less BW was lost from December 7 through calving (linear effect, P = 0.02) as frequency of supplementation increased, but the magnitude of difference in weight change was relatively small. Body condition responded similarly through early February (linear effect, P = 0.02), although treatment effects were not as distinct at calving (cubic effect, P = 0.11). In Exp. 2, 16 ruminally fistulated Hereford x Angus steers (BW = 257 kg) were blocked by weight and assigned to one of the four frequencies of supplementation. Steers were offered tallgrass prairie hay (73.5% NDF, 4.8% CP) ad libitum and were supplemented at a rate (relative to BW) similar to that of the cows in Exp. 1. Increasing frequency of supplementation increased (linear effect, P < or = 0.02) forage OM intake, OM and NDF digestion, and digestible OM intake. However, the most prominent differences in forage OM intake tended (cubic effect, P = 0.07) to occur with the two extreme frequencies of supplementation. In conclusion, forage use was improved with an increased frequency of supplementation, but the impact on performance is not likely to be large unless extreme differences in frequency occur.

Ammonia↗

Impact of different wheat milling by-products in supplements on the forage use and performance of beef cattle consuming low-quality, tallgrass-prairie forage.

Two experiments were conducted to evaluate the impacts on forage use and beef cattle performance of incorporating divergent wheat milling by-products in a 30% CP supplement. The by-products were wheat bran (high fiber) and second clears (high starch). The by-products were added as 1) 100% wheat bran; 2) 67% wheat bran, 33% second clears; or 3) 33% wheat bran, 67% second clears to constitute approximately 47 to 49% of the supplement. In Exp. 1, 90 Hereford x Angus cows (BW = 554 kg) grazing winter, tallgrass-prairie range were fed the supplement treatments (2.27 kg/cow daily) from early December 1997 until calving (average calving date = 3/11/98). Cumulative BW and condition changes from trial initiation through calving were not significantly different among treatments. Similarly, significant treatment effects on cow pregnancy rates as well as calf birth weights, ADG, and ending weights were not evident. In Exp. 2, 16 ruminally fistulated Hereford x Angus steers (BW = 484 kg) were blocked by weight and assigned to one of the same three supplement treatments or to a negative control (forage only). Steers had ad libitum access to tallgrass-prairie hay (76.4% NDF, 3.1% CP) and were fed supplement at the same rate (relative to BW) as the cows in Exp. 1. Forage OM, NDF, and digestible OM intakes were lower (P < 0.01) for the negative control than for supplemented steers but were not significantly different among the supplemented steers. Digestion of OM was lower (P = 0.03) for the negative control than for supplemented steers, although significant treatment differences were not evident among the supplemented groups. Digestion of NDF was not affected (P = 0.49) by treatment. Within the context of the amount of supplemental protein offered, changes in the combination of wheat milling by-products in the supplement did not affect cow performance or intake and digestion of low-quality forage.

Animal Feed↗

Respiration and gas exchange during recovery from exercise in the American alligator.

Low respiratory exchange ratios (R) occur in reptiles. In crocodilians they have been attributed to the loss of respiratory CO2 into the urine and to the immediate exercise-history of an animal. In this study, expired ventilation (V(E)), oxygen consumption (V(O2)), and carbon dioxide excretion (V(CO2)) were measured during recovery from treadmill-exercise in the American alligator. Both V(O2) and V(CO2) decreased exponentially during recovery, but the decrease of V(CO2) was greater than that of V(O2). By 55 min into recovery R fell to 0.3 +/- 0.02 (mean +/- S.E.M). These low values of R may be partially attributable to hyperventilation that occurred during exercise. These data suggest that both hyperventilation during exercise and a metabolic acidosis deplete blood bicarbonate stores, contributing to a low R during recovery. We propose that the right to left cardiac shunt could facilitate restoration of these alkaline reserves and blood pH.

Alligators and Crocodiles↗

Ventilation and gas exchange during treadmill locomotion in the American alligator (Alligator mississippiensis).

A number of anatomical characters of crocodilians appear to be inconsistent with their lifestyle as sit-and-wait predators. To address this paradoxical association of characters further, we measured lung ventilation and respiratory gas exchange during walking in American alligators (Alligator mississippiensis). During exercise, ventilation consisted of low-frequency, large-volume breaths. The alligators hyperventilated severely during walking with respect to their metabolic demands. Air convection requirements were among the highest and estimates of lung P(CO2) were among the lowest known in air-breathing vertebrates. Air convection requirements dropped immediately with cessation of exercise. These observations indicate that the ventilation of alligators is not limited by their locomotor movements. We suggest that the highly specialized ventilatory system of modern crocodilians represents a legacy from cursorial ancestors rather than an adaptation to a lifestyle as amphibious sit-and-wait predators.

Alligators and Crocodiles↗

Pelvic aspiration in the American alligator (Alligator mississippiensis).

The pelvis of crocodilians is highly derived in that the pubic bones are isolated from the acetabulum and are attached to the ischia via moveable joints. We examined the possible role of this unusual morphology in lung ventilation by measuring ventilation, abdominal pressure and the electrical activity of several abdominal and pelvic muscles in the American alligator (Alligator mississippiensis). We found that the activity of two pelvic muscles, the ischiopubis and ischiotruncus muscles, was correlated with inspiration; these muscles rotate the pubes ventrally and thereby increase abdominal volume. During expiration, contraction of the rectus abdominis and transversus abdominis rotates the pubes dorsally. We suggest that this mechanism facilitates diaphragmatic breathing by creating space for caudal displacement of the viscera during inspiration. Because birds also use a dorso-ventral movement of the pelvis to effect ventilation, some form of pelvic aspiration may be plesiomorphic for archosaurs.

Abdomen↗

Circulatory impairment induced by exercise in the lizard Iguana iguana.

Mechanical integration of the cardiac, muscular and ventilatory pumps enables mammals to vary cardiac output over a wide range to match metabolic demands. We have found this integration lacking in a lizard (Iguana iguana) that differs from mammals because blood flow from the caudal body and ventilation are maximal after, rather than during, exercise. Because Iguana iguana are constrained from ventilation during intense locomotion, they appear to be unable to recruit the abdomen and thorax as a pump for venous return. This constraint on simultaneous running and costal breathing arises from their musculoskeletal design, which is similar to that of basal tetrapods, and so a constraint on venous return during exercise may be ancestral for tetrapods. We suggest that mechanical coupling of the pulmonary and cardiac pumps may have been important for the evolution of high-speed locomotor stamina in terrestrial vertebrates.

Animals↗

Gas exchange potential in reptilian lungs: implications for the dinosaur-avian connection.

The theory that birds evolved from a group of small terrestrial theropod dinosaurs has created much controversy. One argument proposed against this theory is that the lungs of early theropods were incapable of sustaining endothermic gas exchange requirements and could not have given rise to the lungs of birds. A reexamination of the comparative physiological and morphological literature combined with a theoretical analysis of gas exchange potential indicates that non-avian lungs would not constrain the gas exchange requirements of early endotherms. Furthermore, our analysis suggests that factors besides diffusive gas exchange were important in the evolution of the distinct morphology of the highly effective avian and mammalian lungs. reserved.

Animals↗

Contribution of gular pumping to lung ventilation in monitor lizards.

A controversial hypothesis has proposed that lizards are subject to a speed-dependent axial constraint that prevents effective lung ventilation during moderate- and high-speed locomotion. This hypothesis has been challenged by results demonstrating that monitor lizards (genus Varanus) experience no axial constraint. Evidence presented here shows that, during locomotion, varanids use a positive pressure gular pump to assist lung ventilation. Disabling the gular pump reveals that the axial constraint is present in varanids but it is masked by gular pumping under normal conditions. These findings support the prediction that the axial constraint may be found in other tetrapods that breathe by costal aspiration and locomote with a lateral undulatory gait.

Animals↗

Evolution of the vertebrate cardio-pulmonary system.

Vertebrate lungs have long been thought to have evolved in fishes largely as an adaptation for life in hypoxic water. This view overlooks the possibility that lungs may have functioned to supply the heart with oxygen and may continue to serve this function in extant fishes. The myocardium of most vertebrates is avascular and obtains oxygen from luminal blood. Because oxygen-rich pulmonary blood mixes with oxygen-poor systemic blood before entering the heart of air-breathing fishes, lung ventilation may supply the myocardium with oxygen and expand aerobic exercise capabilities. Although sustained exercise in tetrapods is facilitated by septation of the heart and the formation of a dual pressure system, a divided cardio-pulmonary system may conflict with myocardial oxygenation because the right side of the heart is isolated from pulmonary oxygen. This may have contributed to the evolution of the coronary circulation.

Animals↗

Effects of different supplemental sugars and starch fed in combination with degradable intake protein on low-quality forage use by beef steers.

Twenty ruminally fistulated steers (Exp. 1, 448 kg and Exp. 2, 450 kg) were used in two consecutive randomized complete block experiments with five treatments in each experiment. The purpose was to evaluate the impact of feeding different supplemental sugars or starch in combination with supplemental degradable intake protein (DIP) on the utilization of low-quality tallgrass-prairie hay. In Exp. 1, steers were given ad libitum access to forage and, except for the negative control (NC), received a limited supply (insufficient to maximize forage use) of supplemental DIP (.031% BW/d, DM basis). In addition to the NC, this experiment included four supplementation treatments in which one of four carbohydrate (CHO) sources (starch, glucose, fructose, or sucrose) was fed at .30% BW of DM/d. In Exp. 2, the treatment structure was identical except that the supplemental DIP level (.122% BW, DM basis) was near the level needed to maximize forage use. Forage OM intake (FOMI) was not affected (P> or =.26) by supplementation in Exp. 1 but was increased (P = .05) in Exp. 2. However, no difference (P> or =.46) in FOMI occurred among CHO sources in either experiment. Total OM and digestible OM intakes were increased (P<.01) by supplementation in both experiments. In Exp. 1, no difference (P>.26) in OM digestion (OMD) occurred among treatments. In Exp. 2, supplementation increased (P<.01) OMD. Additionally, sugars yielded a higher (P = .04) OMD than starch, and the monosaccharides yielded a higher (P = .02) OMD than sucrose. In Exp. 1, NDF digestion (NDFD) was decreased (P = .02) by supplementation, but no differences (P> or =.21) occurred among CHO sources. In Exp. 2, NDFD was increased (P = .03) by supplementation. Additionally, sugars led to higher (P = .05) NDFD than starch, and the monosaccharides led to higher (P = .03) NDFD than sucrose. In both experiments, discernible patterns were observable with regard to the effects of supplementation and type of supplemental CHO on ruminal fermentation characteristics. In conclusion, even though some consistency in fermentation profiles for different carbohydrate sources was evident in both experiments, forage intake and digestion responses were not consistent across experiments. This raises the possibility that carbohydrate source may interact with the amount of supplemental DIP fed and, as such, deserves additional investigation.

Animal Feed↗

Lack of locomotor-cardiac coupling in trotting dogs.

Coupling of locomotor and cardiac cycles has been suggested to facilitate effective arterial delivery and venous return during vigorous exercise. In an attempt to document locomotor-cardiac coupling, we ran five dogs on a motorized treadmill while monitoring heart activity with surface electrocardiogram electrodes and locomotor events with high-speed video and an accelerometer mounted on the dog's back. Analysis of the cardiac and locomotor frequencies revealed that heart rate was usually slightly greater than stride frequency. Hence the timing of the cardiac cycles varied with respect to the phase of the locomotor cycles, and therefore consistent coupling of the locomotor and cardiac cycles was not observed in any of the dogs. However, the period of the cardiac cycle sometimes varied in a rhythmic way that caused brief periods of transient coupling of the locomotor and cardiac cycles in three of the five dogs. These brief periods of coupling (5-20 heartbeats) occurred at approximately the same phase relationship in each of the three dogs. We hypothesize that the variation in cardiac period and the resulting transient coupling are a function of locomotor and ventilatory influences on venous return and/or ventricular ejection. Because venous return and ejection fraction are likely to vary in an unpredictable manner when animals run in a complex environment, we suggest that reflex control of heart rate will be important during locomotion and strict integer coupling of the locomotor and cardiac cycles is unlikely to evolve.

Animals↗